Epinephelus

Epinephelus; Temporal range: 55–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N ; Eocene to present
	Epinephelus fasciatus, the type species
	Epinephelus tukula
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Actinopterygii
Order:	Perciformes
Family:	Serranidae
Subfamily:	Epinephelinae
Tribe:	Epinephelini
Genus:	Epinephelus; Bloch, 1793
Type species
	Epinephelus marginalis; Bloch, 1793
Species
	see text
Synonyms
	Altiserranus Whitley, 1947; Cerna Bonaparte, 1833; Cernua [Costa, 1849; Cynichthys Swainson, 1839; Enneistus D.S. Jordan & Evermann, 1896; Garrupa D.S. Jordan, 1890; Homalogrystes Alleyne & Macleay, 1877; Hyposerranus Klunzinger, 1884; Itaiara Vaillant & Bocourt, 1878; Labroperca Gill, 1862; Merou Bonaparte (ex Cuvier), 1831; Merus Poey (ex Cuvier), 1871; Phrynotitan Gill, 1885; Priacanthichthys Day, 1868; Promicrops Poey (ex Gill), 1868; Stereolepoides Fowler, 1923;

Epinephelus is a genus of marine ray-finned fish, groupers from the subfamily Epinephelinae, part of the family Serranidae, which also includes the anthias and sea basses. They are predatory fish, largely associated with reefs and are found in tropical and subtropical seas throughout the world. They are important target species for fisheries.

Characteristics

The fishes in the genus Epinephelus have elongate, subcylindrical bodies which may be oblong or deep and compressed in shape. The depth of the body varies between 2.3 and 3.7 times the standard length and head is usually around the same length as the body is deep. The preopercle can be rounded or angular and has a serrated rear edge with the serrations at the angle being enlarged to a lesser or greater extent. In a small number of species serrations are small and on the lower edge they are covered by skin. Caniform teeth are found at the front of jaws, although these can be rather small in a few species. They do not have any obviously enlarged caniform teeth in the middle of the lower jaw. There are teeth on the roof of the mouth. In adults, the maxilla does not have a noticeable bony protrusion on the lower rear angle, although they can have a deep step or hook-like process which is hidden by the upper lips, on the rear part of its lower edge. The dorsal fin normally contains 9 spines, although some species have 10, as well as 12 to 19 rays. The origin of the dorsal fin sits above the opercle and the soft rayed part is shorter than the spiny part. The anal fin contains 3 distinct spines and 7 to 10 soft rays. The pectoral fin is rounded with its middle rays being longer than the others longest. The caudal fin may be rounded, truncate or concave, contains 8 branched rays and 8 to 10 fin rays which are slender, unbranched and unsegmented (referred to as "procurrent") fin rays at the leading edges of he caudal fin on the upper lobe and 7 branched rays and 7 to 10 procurrent rays in the lower lobe. The body is covered in ctenoid or smooth scales.^[4]

Habitat and biology

Epinephelus groupers are occur mainly on coral or rocky reefs, although a small number of species have been recorded over substrates consisting of sand, silt or mud. A few species are found in deep water, down to at least 525 metres (1,722 ft), but the majority occur between 10 and 200 metres (33 and 656 ft). The two largest members of the genus, E. itajara and E. lanceolatus, either of which may attain a length in excess of 2 metres (6.6 ft) and a weight greater than 400 kilograms (880 lb) have frequently been recorded in estuaries and harbours. Most of the species in the genus Epinephelus are predatory fish which feed on larger invertebrates, mostly crustaceans, and other fishes taken on or close to the substrate. E. undulosus is an unusual grouper species distinguished by having many, long gill rakers and this species has been reported to feed on pelagic tunicates, at least on occasion. Only a few species have had their reproductive biology studied and many species appear to be protogynous hermaphrodites. However, in some species there are males in the populations which are smaller than some of the females, suggesting a more complex biology and this suggests that some females do not change sex, and that some males may not have a undergone a functional female stage.^[4]

Distribution

Epinephelus species are found around the world in tropical and subtropical seas and oceans. The greatest diversity occurs in the Indo-West Pacific, while 8 species are found in the eastern Pacific, 11 in the western Atlantic Ocean 9 species in the eastern Atlantic Ocean and the Mediterranean.^[4] Four species have entered the Mediterranean Sea from the Red Sea via the Suez Canal as Lessepsian migrants.^[5]

Utilisation

Epinephelus groupers are among the most valuable species exploited by commercial fishes in the world's tropical seas and they fetch some of the highest prices when marketed. They have also been used in aquaculture.^[4]

Species

The 89 recognized species in this genus are:^[6]

Epinephelus adscensionis (Osbeck, 1765)
Epinephelus aeneus (É. Geoffroy Saint-Hilaire, 1817) (white grouper)
Epinephelus akaara (Temminck & Schlegel, 1843) (Hong Kong grouper)
Epinephelus albomarginatus Boulenger, 1903 (white-edged grouper)
Epinephelus amblycephalus (Bleeker, 1857) (banded grouper)
Epinephelus analogus T. N. Gill, 1863 (spotted grouper)
Epinephelus andersoni Boulenger, 1903 (cat-face grouper)
Epinephelus areolatus (Forsskål, 1775) (areolate grouper)
Epinephelus awoara (Temminck & Schlegel, 1843) (yellow grouper)
Epinephelus bilobatus J. E. Randall & G. R. Allen, 1987 (twin-spot grouper)
Epinephelus bleekeri (Vaillant, 1878) (dusky-tail grouper)
Epinephelus bontoides (Bleeker, 1855) (pale-margin grouper)
Epinephelus bruneus Bloch, 1793 (long-tooth grouper)
Epinephelus caninus (Valenciennes, 1843) (dog-tooth grouper)
Epinephelus chabaudi (Castelnau, 1861) (moustache grouper)
Epinephelus chlorocephalus (Valenciennes, 1830) (Tonga grouper)
Epinephelus chlorostigma (Valenciennes, 1828) (brown-spotted grouper)
Epinephelus cifuentesi Lavenberg & Grove, 1993 (Olive grouper)
Epinephelus clippertonensis G. R. Allen & D. R. Robertson, 1999
Epinephelus coeruleopunctatus (Bloch, 1790) (white-spotted grouper)
Epinephelus coioides (F. Hamilton, 1822) (orange-spotted grouper)
Epinephelus corallicola (Valenciennes, 1828) (coral grouper)
Epinephelus costae (Steindachner, 1878) (gold-blotch grouper)
Epinephelus craigi Frable, Tucker & Walker, 2018^[7]
Epinephelus cyanopodus (J. Richardson, 1846) (speckled blue grouper)
Epinephelus daemelii (Günther, 1876) (saddle-tail grouper)
Epinephelus darwinensis J. E. Randall & Heemstra, 1991 (Darwin grouper)
Epinephelus diacanthus (Valenciennes, 1828) (spiny-cheek grouper)
Epinephelus drummondhayi Goode & T. H. Bean, 1878
Epinephelus epistictus (Temminck & Schlegel, 1843) (dotted grouper)
Epinephelus erythrurus (Valenciennes, 1828) (cloudy grouper)
Epinephelus fasciatomaculosus (W. K. H. Peters, 1865) (rock grouper)
Epinephelus fasciatus (Forsskål, 1775) (black-tip grouper)
Epinephelus faveatus (Valenciennes, 1828) (barred-chest grouper)
Epinephelus flavocaeruleus (Lacépède, 1802) (blue-and-yellow grouper)
Epinephelus fuscoguttatus (Forsskål, 1775) (brown-marbled grouper)
Epinephelus fuscomarginatus Johnson & Worthington Wilmer, 2019 ^[8]
Epinephelus gabriellae J. E. Randall & Heemstra, 1991 (multispotted grouper)
Epinephelus geoffroyi (Klunzinger, 1870) (Red Sea spotted grouper)^[9]
Epinephelus goreensis (Valenciennes, 1830) (Dungat grouper)
Epinephelus guttatus (Linnaeus, 1758) (red hind)
Epinephelus heniochus Fowler, 1904 (Bridled grouper)
Epinephelus hexagonatus (Bloch & J. G. Schneider, 1801) (star-spotted grouper)
Epinephelus howlandi (Günther, 1873) (black-saddle grouper)
Epinephelus indistinctus J. E. Randall & Heemstra, 1991 (Somali grouper)
Epinephelus irroratus (Forster, 1801) (Marquesan grouper)
Epinephelus itajara (Lichtenstein, 1822) (Atlantic goliath grouper)
Epinephelus kupangensis Tucker, Kurniasih & Craig, 2016 ^[10]
Epinephelus labriformis (L. Jenyns, 1840) (starry grouper)
Epinephelus lanceolatus (Bloch, 1790) (giant grouper)
Epinephelus latifasciatus (Temminck & Schlegel, 1843) (striped grouper)
Epinephelus lebretonianus (Hombron & Jacquinot, 1853) (mystery grouper)
Epinephelus longispinis (Kner, 1864) (long-spine grouper)
Epinephelus macrospilos (Bleeker, 1855) (snub-nose grouper)
Epinephelus maculatus (Bloch, 1790) (high-fin grouper)
Epinephelus magniscuttis Postel, Fourmanoir & Guézé, 1963 (speckled grouper)
Epinephelus malabaricus (Bloch & J. G. Schneider, 1801) (Malabar grouper)
Epinephelus marginatus (R. T. Lowe, 1834) (dusky grouper)
Epinephelus melanostigma Schultz, 1953 (one-blotch grouper)
Epinephelus merra Bloch, 1793 (honeycomb grouper)
Epinephelus miliaris (Valenciennes, 1830) (net-fin grouper)
Epinephelus morio (Valenciennes, 1828) (red grouper)
Epinephelus morrhua (Valenciennes, 1833) (comet grouper)
Epinephelus multinotatus (W. K. H. Peters, 1876) (white-blotched grouper)
Epinephelus ongus (Bloch, 1790) (white-streaked grouper)
Epinephelus poecilonotus (Temminck & Schlegel, 1843) (dot-dash grouper)
Epinephelus polylepis J. E. Randall & Heemstra, 1991 (small-scaled grouper)
Epinephelus polyphekadion (Bleeker, 1849) (camouflage grouper)
Epinephelus polystigma (Bleeker, 1853) (white-dotted grouper)
Epinephelus posteli Fourmanoir & Crosnier, 1964 (striped-fin grouper)
Epinephelus quinquefasciatus (Bocourt, 1868) (Pacific goliath grouper)
Epinephelus quoyanus (Valenciennes, 1830) (long-fin grouper)
Epinephelus radiatus (F. Day, 1868) (oblique-banded grouper)
Epinephelus retouti Bleeker, 1868 (red-tipped grouper)
Epinephelus rivulatus (Valenciennes, 1830) (halfmoon grouper)
Epinephelus sexfasciatus (Valenciennes, 1828) (six-bar grouper)
Epinephelus socialis (Günther, 1873) (surge grouper)
Epinephelus spilotoceps Schultz, 1953 (four-saddle grouper)
Epinephelus stictus J. E. Randall & G. R. Allen, 1987 (black-dotted grouper)
Epinephelus stoliczkae (F. Day, 1875) (epaulet grouper)
Epinephelus striatus (Bloch, 1792) (Nassau grouper)
Epinephelus suborbitalis Amaoka & J. E. Randall, 1990 (seamount grouper)
Epinephelus summana (Forsskål, 1775) (Summan grouper)
Epinephelus tankahkeei Wu, Qu, Lin, Tang & Ding 2020^[11]
Epinephelus tauvina (Forsskål, 1775) (greasy grouper)
Epinephelus timorensis J. E. Randall & G. R. Allen, 1987 (yellow-spotted grouper)
Epinephelus trimaculatus (Valenciennes, 1828) (three-spot grouper)
Epinephelus trophis J. E. Randall & G. R. Allen, 1987 (plump grouper)
Epinephelus tuamotuensis Fourmanoir, 1971 (reticulated grouper)
Epinephelus tukula Morgans, 1959 (potato grouper)
Epinephelus undulatostriatus (W. K. H. Peters, 1867) (Maori grouper)
Epinephelus undulosus (Quoy & Gaimard, 1824) (wavy-lined grouper)

Some of these species are placed in the genus Hyporthodus by some authorities, for example Epinephelus darwinensis is treated as Hyporthodus darwinensis by the Catalog of Fishes.^[12]